外源H_2O_2和AsA对大蒜试管苗玻璃化及内源活性氧代谢特征的影响
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摘要
大蒜(Allium sativum L.)是重要的药食兼用植物,在世界各地普遍栽培。大蒜栽培种主要利用鳞茎进行无性繁殖,因此极易造成病毒积累。通过组织培养技术脱毒是大蒜品种提纯复壮的有效手段,但是大蒜试管苗培养过程中玻璃化现象严重,因此对玻璃化机理的研究具有重要的理论意义与应用价值。近年来的研究表明,试管苗玻璃化过程中存在不同程度的氧化胁迫。与正常试管苗相比,玻璃化苗的O2·-产生速率和H2O2含量均提高,脂质过氧化程度增强,保护酶调节功能紊乱,说明试管苗玻璃化与活性氧代谢相关。
本试验以大蒜品种‘二水早’为试材,研究了外源H2O2对大蒜试管苗玻璃化及活性氧代谢动态变化的影响,外源H2O2和AsA作用下内源活性氧的产生和定位,以及活性氧清除剂AsA对外源H2O2诱导下大蒜试管苗玻璃化及活性氧代谢动态变化的影响,从而进一步明确活性氧代谢与大蒜试管苗玻璃化的关系,从活性氧代谢的角度探讨大蒜试管苗玻璃化的机理,为试管苗玻璃化的控制提供理论依据。主要研究结果如下：
1.H2O2胁迫影响试管苗生长,2.0 mmol·L-1 H2O2处理显著促进试管苗玻璃化的发生；在H2O2胁迫下,试管苗SOD、POD、CAT和APX活性先升高后降低,MDA和膜透性水平明显提高。说明氧化胁迫可诱导试管苗玻璃化,脂质过氧化可能是试管苗玻璃化的诱因之一。
2.O2·-和H2O2的组织化学染色结果表明,各处理下活性氧产生和积累的区域均主要集中在试管苗叶片和假茎。亚细胞定位结果表明,H2O2主要积累在靠近叶绿体一侧的细胞壁及细胞间隙。外源H2O2胁迫下大蒜试管苗内源活性氧产生和积累量较多,外源H2O2+AsA处理与正常培养条件下内源活性氧产生和积累量较少。
3.添加活性氧清除剂AsA,能诱导外源H2O2胁迫下试管苗保护酶活性增强,抗氧化物质含量提高,O2·-产生速率和内源H2O2含量降低,MDA含量减少,膜透性下降,表明AsA能有效减缓由H2O2胁迫诱导的活性氧的伤害作用,从而显著降低试管苗玻璃化的发生。
Garlic (Allium sativum L.) is an important edible and medicinal plant grown worldwide. Virus-free and rapid propagation is an efficient means in purification and rejuvenation during the production of garlic. But the existence of hyperhydricity in garlic tissue culture is a major problem; therefore the study of hyperhydricity was of great theoretic significance and application value. Previous research showed that there are oxidative stress at different degree in hyperhydricity during in vitro culture, the O2·-generation rate and H2O2 level were significantly higher than that of the normal shoots, the function of protective enzymes of hyperhydric shoots was abnormal and the hyperhydric shoots were stressed and destroyed by oxygen free radicals more heavily than normal shoots, which revealed that hyperhydricity is closely connected with reactivated oxygen species metabolism.
In order to clarify the mechanism of garlic hyperhydricity induced by reactive oxygen species and provide theoretical basis for control the hyperhydricity, in this experiment, using the garlic variety'Ershuizao'as test materials, the effects of hyperhydricity and reactive oxygen species metabolism under exogenous H2O2 stress and the occurrence and location of endogenous reactive oxygen species and the effects of hyperhydricity and reactive oxygen species metabolism of the garlic plantlet in vitro under exogenous H2O2 stress, exogenous 'H2O2+AsA' treatment and normal culture conditions were studied. The main results are as follows:
1. Exogenous H2O2 influenced the normal growth of plantlet and 2.0 mmol·L-1 H2O2 stress significantly enhanced the hyperhydricity of the garlic plantlet in vitro. The activity of SOD, POD, CAT and APX increased at first and then decreased, the MDA levels and the rate of elect rolyte leakage was increased under oxidative stress during the culture. The result indicated that the oxygen free radicals can induce the occurrence of hyperhydricity and the lipid peroxidation may be one of the induction factors of hyperhydricity.
2. The histochemical localization of O2·- and H2O2 demonstrated that endogenous reactive oxygen species accumulated mainly in leaves and pesudostem under different treatments. Cytochemical staining showed that H2O2 accumulation was observed mostly in cell walls and intercellular space nearby chloroplast. The studies preliminarily confirmed that exogenous H2O2 stress enhanced the production of endogenous O2·- and H2O2 of the garlic plantlet in vitro effectively and application of exogenous AsA significantly decreased the accumulation of reactive oxygen species induced by H2O2 stress.
3. The results showed that application of exogenous AsA increased the activity of protective enzymes and the contents of antioxidant substances, and decreased the level of lipid peroxidation and the accumulation of reactive oxygen species of the garlic plantlet in vitro under exogenous H2O2 stress,. There was certain alleviate effects of exogenous AsA on the damage of reactive oxygen species induced by H2O2 stress of the the garlic plantlet in vitro, thereby reduce the hyperhydricity plantlet rates significantly.
本试验以大蒜品种‘二水早’为试材,研究了外源H2O2对大蒜试管苗玻璃化及活性氧代谢动态变化的影响,外源H2O2和AsA作用下内源活性氧的产生和定位,以及活性氧清除剂AsA对外源H2O2诱导下大蒜试管苗玻璃化及活性氧代谢动态变化的影响,从而进一步明确活性氧代谢与大蒜试管苗玻璃化的关系,从活性氧代谢的角度探讨大蒜试管苗玻璃化的机理,为试管苗玻璃化的控制提供理论依据。主要研究结果如下：
1.H2O2胁迫影响试管苗生长,2.0 mmol·L-1 H2O2处理显著促进试管苗玻璃化的发生；在H2O2胁迫下,试管苗SOD、POD、CAT和APX活性先升高后降低,MDA和膜透性水平明显提高。说明氧化胁迫可诱导试管苗玻璃化,脂质过氧化可能是试管苗玻璃化的诱因之一。
2.O2·-和H2O2的组织化学染色结果表明,各处理下活性氧产生和积累的区域均主要集中在试管苗叶片和假茎。亚细胞定位结果表明,H2O2主要积累在靠近叶绿体一侧的细胞壁及细胞间隙。外源H2O2胁迫下大蒜试管苗内源活性氧产生和积累量较多,外源H2O2+AsA处理与正常培养条件下内源活性氧产生和积累量较少。
3.添加活性氧清除剂AsA,能诱导外源H2O2胁迫下试管苗保护酶活性增强,抗氧化物质含量提高,O2·-产生速率和内源H2O2含量降低,MDA含量减少,膜透性下降,表明AsA能有效减缓由H2O2胁迫诱导的活性氧的伤害作用,从而显著降低试管苗玻璃化的发生。
Garlic (Allium sativum L.) is an important edible and medicinal plant grown worldwide. Virus-free and rapid propagation is an efficient means in purification and rejuvenation during the production of garlic. But the existence of hyperhydricity in garlic tissue culture is a major problem; therefore the study of hyperhydricity was of great theoretic significance and application value. Previous research showed that there are oxidative stress at different degree in hyperhydricity during in vitro culture, the O2·-generation rate and H2O2 level were significantly higher than that of the normal shoots, the function of protective enzymes of hyperhydric shoots was abnormal and the hyperhydric shoots were stressed and destroyed by oxygen free radicals more heavily than normal shoots, which revealed that hyperhydricity is closely connected with reactivated oxygen species metabolism.
In order to clarify the mechanism of garlic hyperhydricity induced by reactive oxygen species and provide theoretical basis for control the hyperhydricity, in this experiment, using the garlic variety'Ershuizao'as test materials, the effects of hyperhydricity and reactive oxygen species metabolism under exogenous H2O2 stress and the occurrence and location of endogenous reactive oxygen species and the effects of hyperhydricity and reactive oxygen species metabolism of the garlic plantlet in vitro under exogenous H2O2 stress, exogenous 'H2O2+AsA' treatment and normal culture conditions were studied. The main results are as follows:
1. Exogenous H2O2 influenced the normal growth of plantlet and 2.0 mmol·L-1 H2O2 stress significantly enhanced the hyperhydricity of the garlic plantlet in vitro. The activity of SOD, POD, CAT and APX increased at first and then decreased, the MDA levels and the rate of elect rolyte leakage was increased under oxidative stress during the culture. The result indicated that the oxygen free radicals can induce the occurrence of hyperhydricity and the lipid peroxidation may be one of the induction factors of hyperhydricity.
2. The histochemical localization of O2·- and H2O2 demonstrated that endogenous reactive oxygen species accumulated mainly in leaves and pesudostem under different treatments. Cytochemical staining showed that H2O2 accumulation was observed mostly in cell walls and intercellular space nearby chloroplast. The studies preliminarily confirmed that exogenous H2O2 stress enhanced the production of endogenous O2·- and H2O2 of the garlic plantlet in vitro effectively and application of exogenous AsA significantly decreased the accumulation of reactive oxygen species induced by H2O2 stress.
3. The results showed that application of exogenous AsA increased the activity of protective enzymes and the contents of antioxidant substances, and decreased the level of lipid peroxidation and the accumulation of reactive oxygen species of the garlic plantlet in vitro under exogenous H2O2 stress,. There was certain alleviate effects of exogenous AsA on the damage of reactive oxygen species induced by H2O2 stress of the the garlic plantlet in vitro, thereby reduce the hyperhydricity plantlet rates significantly.
引文
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